Oryctes rhinoceros Scientific Name Oryctes rhinoceros (Linnaeus, 1758) Synonyms: Oryctes stentor Castelnau, 1840 Scarabaeus rhinoceros Linnaeus Common Name Coconut rhinoceros beetle (CRB), coconut beetle, rhinoceros beetle, palm rhinoceros beetle, Asiatic rhinoceros beetle Figure 1. Oryctes rhinoceros female (Mark Schmaedick, American Samoa Community College). Type of Pest Beetle Taxonomic Position Class: Insecta, Order: Coleoptera, Family: Scarabaeidae Reason for Inclusion in Manual Figure 2. Oryctes rhinoceros male (Aubrey Moore, Suggestion from CAPS Community University of Guam). Pest Description Descriptions of all life cycles can be found in Gressitt (1953). Eggs: Eggs are whitish brown and 3 to 1 3 4 mm (approx. /8 to /16 in) (Giblin- Davis, 2001). Eggs are initially soft and oblong but swell into a rubbery circle four to five days after being laid (Hinckley, 1973). Larvae: Newly hatched larvae are 7.5 5 mm long (approx. /16 in) (Lever, 1979). “The large (60 to 105 mm long [approx. 3 1 2 /8 to 4 /8 in]) white mature larva is C- shaped, with a brown head capsule and legs. The posterior part of the abdomen is a bluish-grey colour” Figure 3. Larva of Oryctes rhinoceros (Mark (Giblin-Davis, 2001). Larval instars are Schmaedick, American Samoa Community College). Last Updated: February 20, 2014 1 differentiated by the size of their head capsules (Gressitt, 1953). Pupae: “The color is somewhat yellowish brown and the body has a somewhat rubbery appearance. Most of the external adult structures can be detected…the pupa ranges 9 3 15 from 39.4 to 51.5 mm. [approx. 1 /16 to 2 in] long, 19-23.6 [approx. /4 to /16 in] wide, 5 3 and 16 to 19.4 [approx. /8 to /4 in] deep…the surface is rather smooth and shiny, with minute punctures or bristles in part. The ventral surfaces and legs bear distinct reddish bristles. The sclerotization of the exoskeleton is highly developed, and the beetle is extremely strong. The sides of the mandibles and parts of the legs and ventral surfaces of the body are clothed with bright reddish brown hairs” (Gressitt, 1953). 3 1 Adults: Adults range from 30 to 57 mm (approx. 1 /16 to 2 /4 in) (Giblin-Davis, 2001) and are black “with a fine reddish pubescence on the ventral surface.” Adults “bear on the head a median horn curved backward; the horn is much longer in the male than in the female” (Lever, 1979). Male: “The male has a large horn on its head and a smooth and shiny abdomen” (Vargo, 1995). Female: “The female has a smaller horn [than the male] but is distinguished by long, erect, reddish hairs that are visible at the tip of the abdomen” (Vargo, 1995). There can be considerable variation in body size and the length of the cephalic horn in both males and females. This is correlated with the larval environment (Gressitt, 1953). Biology and Ecology The life cycle can range from 4 to 9 months. More than one generation can occur per year (Giblin-Davis, 2001); when conditions are favorable there may be more than 3 generations per year. All stages of the pest may be found at the same time (Gressitt, 1953). Adults mate several times in dead or dying plant material (Catley, 1969), including decaying palm trunks (Zelazny and Alfiler, 1987). The females make a serpentine burrow in which they lay their eggs. According to Hinckley (1973), females can oviposit in “almost any log or heap soft enough for burrowing, yet firm enough to provide compacted frass”. Females lay between 3 and 4 Figure 4. Breeding site of Oryctes rhinoceros under a clutches of eggs with about 30 eggs coconut log (Mark Schmaedick, American Samoa per clutch (Hinckley, 1973), laying Community College). between 70 and 140 eggs total Last Updated: February 20, 2014 2 (Giblin-Davis, 2001). After oviposition, they cover them with the chewed material (Hinckley, 1973). Males usually arrive after the females lay their eggs and stay longer in the breeding site to prepare the wood for the larvae (Zelazny and Alfiler, 1991; Alfiler, 1999). Zelazny and Alifiler (1987) described four phases of the adult life of O. rhinoceros in the Philippines: immature phase, first feeding, breeding phase, and late life feeding. The last two phases occurred simultaneously. In late life feeding, the beetles will occasionally make additional visits to palms (reviewed in Alfiler, 1999). Eggs are laid in manure, rotten wood, decaying vegetation found in coconut plantations, or other organic matter (Giblin-Davis, 2001; Schmaedick, 2005). Trees damaged by typhoons can serve as breeding sites (Moore, 2007). Bedford (1976) states that dead standing palms are the most favored breeding sites. Eggs hatch in about 12 days (Giblin-Davis, 2001). Larvae can be found in decaying or dead standing palms (Giblin-Davis, 2001) and eat rotting organic matter (Muniappan, 2002). They may be found feeding in the trunks of living palms, but this is only when the tree is already seriously injured by the adult or other means (Gressitt, 1953). In Guam, a significant number of O. rhinoceros have been found to complete their entire life cycle in the crowns of coconut palms with larvae feeding on decaying detritus caught between the petioles (Moore, 2012). Crowns are only damaged by adults as they bore into folded, emerging fronds to feed on sap. In lab conditions, larvae preferred temperatures around 27 to 29°C (81° to 84°F) and had a strong negative phototaxis response, possibly an adaptation against desiccation and predation (Bedford, 1980). They avoid areas where temperatures are above 37°C (100°F) (Hinckley, 1973). Larvae seem to prefer areas with high moisture content, with more larvae being found in material when moisture content was 80 to 100% (Kamarudin et al., 2005). Earlier work found a moisture content over 77% was needed for satisfactory larval development (reviewed in Kamarudin et al., 2005). They develop fully in 72 to 130 days (Giblin-Davis, 2001) going through three instars (Gressitt, 1953; Hinckley, 1973). Once mature, larvae go through a nonfeeding pre-pupal stage lasting 8 to 13 days before pupation occurs (Schmaedick, 2005). Pupation usually occurs in the soil but can also occur in the larval habitats as well (Giblin-Davis, 2001). Larvae construct oval pupal cells measuring 55 x 35 x 33 mm 3 3 5 (approx. 2 /16 x 1 /8 x 1 /16 in) enclosing it with sawdust and frass (Gressitt, 1953) and sometimes construct cocoons of soil or plant tissue when pupating in softer material (Vargo, 1995). Pupation lasts about 20 days. Adults will remain in the cocoons after pupation is complete for about 11 to 20 days (Giblin-Davis, 2001). This is to allow the exoskeleton time to darken and harden (Hinckley, 1973). Last Updated: February 20, 2014 3 After emergence, adults remain mostly around the breeding sites (Giblin-Davis, 2001). Adults can fly to the crowns of host plants to feed, flying between dusk and dawn (Catley, 1969). The depth of burrows made by adults can range from 2 to 50 cm 3 11 1 (approx. /4 to 19 /16 in), averaging around 21 cm (approx. 8 /4 in). Adults stay to feed in the burrows several days (Hinckley, 1973). Under favorable conditions, adults are likely to fly short distances whereas longer flights occur when conditions are adverse (Catley, 1969). Hinckley (1973) found that after feeding, adults could travel distances between 2 and 4 km (1.24 to 2.48 mi). Flight is inhibited by rain and moonlight (Vargo, 1995). Adults live for about 3 months (Schmaedick, 2005). Damage Damage is caused by the adult stage, specifically young adults that bore into the crowns of healthy palms. Adults can penetrate 15 11 10 to 50 cm (approx. 3 /16 to 19 /16 in) down close to the center of the spear cluster (Giblin- Davis, 2001). Adults bite through the tightly packed unopened leaves in the central bud. The leaf midribs are most often injured (Lever, 1979). After entering the palm, the beetle feeds on juice produced by host tissue (Giblin-Davis, 2001). It then bores outwards emerging from the base of a central frond (Barber et al., 1971). Figure 5. Palm damage caused by O. Adult feeding damages the inflorescences and rhinoceros adults (Ben Quichocho, USDA- APHIS, Guam). fronds which may decrease or delay fruit production. Mature fronds will often have patches of missing foliage if attacked while in the developing stage (Giblin- Davis, 2001). Damage can appear as V-shaped cuts in the fronds or holes in the midrib (Schmaedick, 2005). Plants killed by O. rhinoceros provide additional breeding sites for future generations (Moore, 2007). There is a positive correlation between abundance of larval food and damage by adults to Figure 6. Feeding damage caused by Oryctes rhinoceros palms. Outbreaks can be (Ernie Nelson, Greenscapes Inc, Guam). Last Updated: February 20, 2014 4 caused by natural catastrophes (typhoons, diseases) or human destruction (replanting, milling, composting, military devastation) (Hinckley, 1973). In older healthy palms, beetle damage can be tolerated. But in palms from 1 to 3 years old, damage by O. rhinoceros can result in severe malformation or death (Giblin-Davis, 2001). Death of the palm occurs when the beetle bores down to the meristematic tissue which can destroy the growing point. This is more likely to occur in young palms with small crowns (Catley, 1969). Attacks are typically concentrated on the margins of palm groves and on taller palms (reviewed in Bedford, 1980). Adults chew the plant tissue to extract the juices and leave a fibrous frass which is pushed out of the burrow entrance. This extruded material is a sign that adults are still present in the host (Catley, 1969; Giblin-Davis, 2001).